Optimal foraging and feeding mode shifts in fishes

Synopsis Most optimal foraging models for fishes are based on particulate feeding behavior. But many obligate planktivores also filter zooplankton. I suggest that feeding mode shifts (e.g. from particulate feeding to filtering) may be predictable from the costs and benefits of foraging in various modes. Quantitative examples of feeding mode shifts in three species of fishes (northern anchovy, pacific mackerel and alewife) from 3 different families support this hypothesis. Feeding mode shifts seem to depend on relative profitability of each mode, but improvements in model predictions will need to include the effects of spatial and temporal patchiness on encounter rates of prey of various sizes.     

Study on foraging mechanism of leeches with different feeding habits based on chemoreception and foraging behavior

The leeches Whitmania pigra and Hirudo nipponia live in similar environments but have different feeding habits. At present, there are few studies of the foraging mechanism of leeches with different feeding habits. In this study, we first used maze tests to show that these two species of leeches could locate and distinguish their prey through chemosensory activity without mechanical stimulation. However, the two leech species have different foraging behaviors: Individuals of W. pigra move slowly and repeatedly adjust direction through probing and crawling to detect the location of prey (snails), whereas individuals of H. nipponia move quickly, and after determining the location of food (porcine blood), they quickly swim or crawl to the vicinity of their prey. Scanning electron microscopy (SEM) revealed that there are two types of sensory cilia and pore structures related to mucus secretion in the heads of both leeches. There are two differently sized types of chemoreceptors on the dorsal lip in W. pigra, which may have different functions during foraging, whereas in H. nipponia there is only one type of chemoreceptor, which is small. We detected the chemical components in the natural food of these two leech species by UHPLC–MS. There were 934 metabolites in the body fluid of snails and 751 metabolites in porcine serum; five metabolites unique to the body fluid of snails and to porcine serum were screened as candidate feeding attractants. Of these metabolites, betaine and arginine effectively attracted individuals of W. pigra and H. nipponia, respectively. In summary, leeches with different feeding habits use chemoreceptors to sense external chemical signals when foraging, and there are significant differences between species in foraging behavior, chemoreceptors, and attractants.     

Scatter-hoarding rodents use different foraging strategies for seeds from different plant species

Seed predation and dispersal by scatter-hoarding rodents are key processes that determine seed survival, and thus, plant regeneration within forests. For decades, there has been much debate on the important effects of seed size (one of the most important seed traits) on rodent foraging preference. Furthermore, the possible selective forces in the evolution of seed size may be influenced by primary selectivity and how rodents treat seeds after harvesting. In this study, different-sized seeds from four species (Pinus armandii, Pinus densata, Abies sp., and Viburnum sp.) harvested by scatter-hoarding rodents were studied in an alpine forest in Southwestern China for two consecutive years. Our results showed that seed size influenced rodent foraging preferences, with bigger seeds being preferred over smaller seeds, within and across species. Rodents only removed and cached the larger seeds of P. armandii, and ate the seeds of the other three species in situ. Rodents are purely seed predators for these three species. For the cached seeds of P. armandii, significantly positive correlations were observed between seed size and dispersal distance among both primary and secondary cached seeds in 2006, but not in 2005. Our results indicate that among many coexisting species with widely different-sized seeds, scatter-hoarding rodents played important roles in the seed dispersal of the big-seeded species alone. This caching behavior could offset the limited seed dispersal of large-seeded and wingless species (P. armandii), in comparison with that of small winged seed species (P. densata and Abies sp.) and frugivore-dispersed species (Viburnum sp.).     

Root traits explain different foraging strategies between resprouting life histories

Drought and fire are prevalent disturbances in Mediterranean ecosystems. Plant species able to regrow after severe disturbances (i.e. resprouter life history) have higher allocation to roots and higher water potential during the dry season than coexisting non-resprouting species. However, seedlings of non-resprouters have a higher survival rate after summer drought. We predict that, to counteract their shallow-rooting systems and to maximize seedling survival, non-resprouters have root traits that confer higher efficiency in soil resource acquisition than resprouters. We tested this prediction in seedlings of less than 1.5 months old. We select 13 coexisting woody species (including both resprouters and non-resprouters), grew them in a common garden and measured the following root traits: length, surface, average diameter, root tissue density (RTD), specific root length (SRL), surface:volume ratio (SVR), specific tip density (STD), tip distribution in depth, internal links ratio (ILR), and degree of branching. These root traits were compared between the two resprouting life histories using both standard cross-species and phylogenetic-informed analysis. Non-resprouters showed higher SRL and longer, thinner and more branched laterals, especially in the upper soil layers. The external links (i.e. the most absorptive root region) were also more abundant, longer, thinner and with higher SVR for non-resprouters. The results were supported by the phylogenetic-informed analysis for the root traits most strongly related to soil resource acquisition (SRL, SVR and branching pattern). The seedling root structure of non-resprouters species allows them to more efficiently explore the upper soil layer, whereas seedling roots of resprouters will permit both carbon storage and deep soil penetration.     

Optimal feeding strategies by adaptive mesh selection for fed-batch bioprocesses

The objective of this contribution is the design of optimal feeding strategies for fed-batch bioprocesses, where complex dynamic models with input and state constraints are present. For the solution of this dynamic optimization problem a transformation to a finite dimensional optimization problem is made using piecewise linear control profiles. The optimization of these profiles is performed by a sequential approach, that includes an ODE solver for the solution of the model ODE's. Further an adaptive mesh selection algorithm was investigated for an appropriate discretization of the control profiles. The implementation of the resulting optimal feeding profiles is shown for a process example, namely the production of nikkomycin by Streptomyces tendae. This implementation uses a hierarchical process control framework, that consists of components for process monitoring, state estimation, and trajectory control.     

Spatial and reversal learning in congeneric lizards with different foraging strategies

Environmental demands that require intensive search for mates, food and nest sites are correlated with efficient spatial memory in many mammalian and avian species. This convergence of evidence has led to the view that spatial memory, and the neurological structures associated with it, have been selected in niches that require memory for the location of goal objects. Whether such evolutionary demands are also correlated with nonspatial abilities that require flexible use of associations similar to those required for spatial memory has not been well studied. In addition, correlations between niche types and the use of spatial or nonspatial memory have not been investigated in nonmammalian, nonavian taxa. In this study, we investigated the relationship between foraging strategies and performance on two tasks, one spatial and the other nonspatial, in congeneric lizard species: Acanthodactylus boskianus, an active forager that collects clumped sedentary prey, Acanthodactylus scutellatus, a sit-and-wait predator that collects distributed mobile prey. The two species did not differ in their performance of a spatial memory task, but A. boskianus, the active forager, performed better on the reversal of a visual discrimination, a nonspatial task. These findings question the generality of the spatial adaptation model for vertebrates. We present the pliancy hypothesis, which we developed to account for these results. Copyright 1999 The Association for the Study of Animal Behaviour.     

Generalist versus specialist strategies of plasticity: snail responses to predators with different foraging modes

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Optimal foraging and the traveling salesman

Optimal foraging models are examined that assume animals forage for discrete point resources on a plane and attempt to minimize their travel distance between resources. This problem is similar to the well-known traveling salesman problem: A salesman must choose the shortest path from his home office to all cities on his itinerary and back to his home office again. The traveling salesman problem is in a class of enigmatic problems, called NP-complete, which can be so difficult to solve that animals might be incapable of finding the best solution. Two major results of this analysis are: (1) The simple foraging strategy of always moving to the closest resource site does surprisingly well. More sophisticated strategies of “looking ahead” a small number of steps, choosing the shortest path, then taking a step, do worse if all the resource sites are visited, but do slightly better (less than 10%) if not all the resource sites are visited. (2) Short cyclical foraging routes resulted when resources were allowed to renew. This is suggested as an alternative explanation for “trap-lining” in animals that forage for discrete, widely separated resources.     

Optimal foraging: a bird in the hand released

Optimal foraging theory aims to elucidate strategies that maximize resource intake. Although traditionally used to understand animal foraging behavior, recent evolutionary experiments with viruses offer a new twist on an old idea.     

Optimal foraging: A case for random movement

Summary The bumblebee, Bombus flavifrons, forages randomly with respect to direction on Polemonium foliosissimum. This foraging pattern is as predicted for a system where there is a low probability of revisiting any given flower upon returning to a patch. This low revisitation probability is a function of the floral resource arrangement. It is further shown that B. flavifrons is using the resource distribution to direct its movements. A large percentage of all movements are to nearest neighbors with maximal foraging efficiency gained through minimization of flight distances.     

Optimal foraging in bumblebees: Hunting by expectation

The foraging behaviour literature contains three hypotheses concerned with hunting by expectation. These suggest possible rules animals use to decide when to leave particular feeding sites and search in other places for food. Predictions of the three hypotheses were tested experimentally by varying the quality of plants (amount and distribution of nectar) encountered by bumblebees (Bombus appositus). Results support only a rate expectation hypothesis. Bees left multiflowered plants when the amount of nectar found in the first flower was below a threshold volume. Bees stayed on plants if they received greater than the threshold volume. This threshold nectar volume is close to the amount predicted if a bee forages to maximize its rate of net energy intake.     

Optimal foraging: movement patterns of bumblebees between inflorescences

Nectar-collecting bumblebees are hypothesized to employ rules of movement which result in the maximum net rate of energy gain (i.e., are optimal). The optimal movement rules are derived from a mathematical model and are used to generate predicted patterns of movement. The predicted patterns are compared with field observations. These observations support the hypothesis. An important component of the mathematical model is the memory of the foraging animal. The field data have implications concerning the memory capabilities of the bumblebees.     

Optimal foraging by zooplankton within patches: the case of Daphnia

The motions of many physical particles as well as living creatures are mediated by random influences or 'noise'. One might expect that over evolutionary time scales internal random processes found in living systems display characteristics that maximize fitness. Here we focus on animal random search strategies [G.M. Viswanathan, S.V. Buldyrev, S. Havlin, M.G.E. Da Luz, E.P. Raposo, H.E. Stanley, Optimizing the success of random searches, Nature 401 (1999) 911-914; F. Bartumeus, J. Catalan, U.L. Fulco, M.L. Lyra, G.M. Viswanathan, Optimizing the encounter rate in biological interactions: Lévy versus Brownian stratagies, Phys. Rev. Lett. 88 (2002) 097901 and 89 (2002) 109902], and we describe experiments with the following Daphnia species: D. magna, D. galeata, D. lumholtzi, D. pulicaria, and D. pulex. We observe that the animals, while foraging for food, choose turning angles from distributions that can be described by exponential functions with a range of widths. This observation leads us to speculate and test the notion that this characteristic distribution of turning angles evolved in order to enhance survival. In the case of theoretical agents, some form of randomness is often introduced into search algorithms, especially when information regarding the sought object(s) is incomplete or even misleading. In the case of living animals, many studies have focused on search strategies that involve randomness [H.C. Berg, Random Walks in Biology, Princeton University, Princeton, New Jersey, 1993; A. Okubo, S.A. Levin (Eds.), Diffusion and Ecological Problems: Modern Perspectives, second ed., Springer, New York, 2001]. A simple theory based on stochastic differential equations of the motion backed up by a simulation shows that the collection of material (information, energy, food, supplies, etc.) by an agent executing Brownian-type hopping motions is optimized while foraging for a finite time in a supply patch of limited spatial size if the agent chooses turning angles taken from an exponential distribution with a specific stochastic intensity or 'noise width'. Search strategies that lead to optimization is a topic of high current interest across many disciplines [D. Wolpert, W. MacReady, No free lunch theorems for optimization, IEEE Transactions on Evolutionary Computation 1 (1997) 67].     

Growth and survival of tench larvae fed under different feeding strategies

Two feeding experiments were conducted to evaluate the growth and survival rates of tench Tinca tinca L. larvae when initially fed with a combination of three different brands of Artemia nauplii under two conditions: (A) in the laboratory and (B) on a commercial farm. At the same time, a protozoan culture of the freshwater ciliate Colpoda cucullus was additionally tested in one of the experiments to possibly enhance the initial hunting behaviour of the larvae. The larvae were fed every 4 h from the onset of exogenous feeding up to 14 days of age. Three types of commercial Artemia products, mainly differing in size and high unsaturated fatty acid (HUFA) content, were used. One group was fed with C. cucullus as a starter. The different combinations of Artemia nauplii were used to evaluate possible effects on larval growth. The final growth at 26.3 °C, expressed in length and weight, did not show significant differences, suggesting the use of the most economically feasible Artemia strain studied. The experiments confirmed that using smaller prey during the first 2 days of feeding increases their survival rate, although the mean final survival rate was high (89%). In the experiment carried out at the commercial fish farm facilities, experimental groups were also fed with Artemia nauplii, using the EG type either enriched or not enriched with HUFA. Again, one of the groups was offered the ciliate C. cucullus as initial feed. Final growth showed significant differences when using Colpoda culture as a starter feed, although this test resulted in the lowest survival rate (69%), indicating that further studies on the management of its culture should be undertaken to improve the applicability of the technique. The mean final survival rate was 83%.     

Optimal foraging: Random movement by pollen collecting bumblebees

Summary Two bumblebee species, Bombus bifarius and B. flavifrons, forage randomly with respect to direction when gathering pollen on Potentilla gracilis. Bees avoid revisiting flowers by being able to differentiate recently visited from unvisited flowers. This recognition occurs while bees are flying over open flowers and appears to be a response to the amount of available pollen within flowers. Random foraging with respect to direction is the optimal strategy when the probability of flower revisitation is low. Bumblebees appear to be moving preferentially between nearest neighbors, again as predicted by foraging theory. This behavior causes the establishment of pollen patches in the P. gracilis population. Unlike other pollinators studied in similar situations, bumblebees on P. gracilis do not forage utilizing an area-restricted searching behavior. Because floral reward quality can be assessed at low cost by bees foraging on P. gracilis, their tendency to move to nearby flowers even after encountering a poor quality blossom apparently yields a higher rate of net energy intake than does area-restricted searching. The data indicate that bumblebees exhibit great plasticity in foraging behavior and that they are able to forage efficiently under a wide range of environmental conditions.     

Optimal foraging in patches: A case for stochasticity

Like much mathematical modeling in biology, most optimal foraging theory is developed from deterministic analogs of basically stochastic processes. Unlike other models, however, it cannot depend on laws of large numbers to justify this simplification; ignoring stochasticity can lead to wrong answers. This is demonstrated for a predator searching spatially separated patches of prey; it is shown that the choice of an optimal procedure for deciding when to leave a patch must be based on a stochastic model—a predator whose procedure is based on a deterministic model can do arbitrarily badly by comparison with the stochastic optimizer. A general solution is given, and its complexity suggests some objections to standard optimality arguments, and some possible alternatives.     

Optimal foraging: food patch depletion by ruddy ducks

Summary I studied the foraging behavior of ruddy ducks (Oxyura jamaicensis) feeding on patchily distributed prey in a large (5-m long, 2-m wide, and up to 2-m deep) aquarium. The substrate consisted of a 4x4 array of wooden trays (1.0-m long, 0.5-m wide, and 0.1-m deep) which contained 6 cm of sand. Any tray could be removed from the aquarium and loaded with a known number of prey. One bird foraged in the aquarium at a time; thus, by removing a food tray after a trial ended and counting the remaining prey, I calculated the number of prey consumed by the bird. I designed several experiments to determine if ruddy ducks abandoned a food patch in a manner consistent with the predictions of a simple, deterministic, patch depletion model. This model is based on the premise that a predator should maximize its rate of net energy intake while foraging. To accomplish this, a predator should only remain in a food patch as long as its rate of energy intake from that patch exceeds the average rate of intake from the environment. In the majority of comparisons, the number of food items consumed by the ruddy ducks in these experiments was consistent with the predictions of the foraging model. When the birds did not forage as predicted by the model, they stayed in the patch longer and consumed more prey than predicted by the model. An examination of the relation between rate of net energy intake and time spent foraging in the food patch indicated that by staying in a patch longer than predicted, the ruddy ducks experienced only a small deviation from maximum rate of net energy intake. These results provided quantitative support for the prediction that ruddy ducks maximize their rate of net energy intake while foraging.     

Optimal foraging area: Size and allocation of search effort

A model is derived for the optimal spatial allocation of foraging effort for an animal returning with food to a central place in a uniform habitat. The forager is assumed to maximize its yield of food during a given period. Foraging effort is expended on search for food, and on transportation to the central place. It is shown that the allocation of search has been optimal if and only if the “marginal cost” of additional food is equal throughout the foraging area when the period has elapsed. The model is used to predict the optimal area radius and allocation of search time. With realistic parameter values, the optimal time per unit area roughly decreases linearly with the distance from the central place. The influence of food density and forager characteristics is examined.